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Census of Antarctic Marine Life SCAR-Marine Biodiversity Information Network BIOGEOGRAPHIC ATLAS OF THE SOUTHERN OCEAN CHAPTER 3.3. PALAEO-OCEANOGRAPHY (BOX). Gersonde R., 2014. In: De Broyer C., Koubbi P., Griffiths H.J., Raymond B., Udekem d’Acoz C. d’, et al. (eds.). Biogeographic Atlas of the Southern Ocean. Scientific Committee on Antarctic Research, Cambridge, pp. 43. EDITED BY: Claude DE BROYER & Philippe KOUBBI (chief editors) with Huw GRIFFITHS, Ben RAYMOND, Cédric d’UDEKEM d’ACOZ, Anton VAN DE PUTTE, Bruno DANIS, Bruno DAVID, Susie GRANT, Julian GUTT, Christoph HELD, Graham HOSIE, Falk HUETTMANN, Alexandra POST & Yan ROPERT-COUDERT SCIENTIFIC COMMITTEE ON ANTARCTIC RESEARCH THE BIOGEOGRAPHIC ATLAS OF THE SOUTHERN OCEAN The “Biogeographic Atlas of the Southern Ocean” is a legacy of the International Polar Year 2007-2009 (www.ipy.org) and of the Census of Marine Life 2000-2010 (www.coml.org), contributed by the Census of Antarctic Marine Life (www.caml.aq) and the SCAR Marine Biodiversity Information Network (www.scarmarbin.be; www.biodiversity.aq). The “Biogeographic Atlas” is a contribution to the SCAR programmes Ant-ECO (State of the Antarctic Ecosystem) and AnT-ERA (Antarctic Thresholds- Ecosystem Resilience and Adaptation) (www.scar.org/science-themes/ecosystems). Edited by: Claude De Broyer (Royal Belgian Institute of Natural Sciences, Brussels) Philippe Koubbi (Université Pierre et Marie Curie, Paris) Huw Griffiths (British Antarctic Survey, Cambridge) Ben Raymond (Australian Antarctic Division, Hobart) Cédric d’Udekem d’Acoz (Royal Belgian Institute of Natural Sciences, Brussels) Anton Van de Putte (Royal Belgian Institute of Natural Sciences, Brussels) Bruno Danis (Université Libre de Bruxelles, Brussels) Bruno David (Université de Bourgogne, Dijon) Susie Grant (British Antarctic Survey, Cambridge) Julian Gutt (Alfred Wegener Institute, Helmoltz Centre for Polar and Marine Research, Bremerhaven) Christoph Held (Alfred Wegener Institute, Helmoltz Centre for Polar and Marine Research, Bremerhaven) Graham Hosie (Australian Antarctic Division, Hobart) Falk Huettmann (University of Alaska, Fairbanks) Alix Post (Geoscience Australia, Canberra) Yan Ropert-Coudert (Institut Pluridisciplinaire Hubert Currien, Strasbourg) Published by: The Scientific Committee on Antarctic Research, Scott Polar Research Institute, Lensfield Road, Cambridge, CB2 1ER, United Kingdom (www.scar.org). Publication funded by: - The Census of Marine Life (Albert P. Sloan Foundation, New York) - The TOTAL Foundation, Paris. The “Biogeographic Atlas of the Southern Ocean” shared the Cosmos Prize awarded to the Census of Marine Life by the International Osaka Expo’90 Commemorative Foundation, Tokyo, Japan. Publication supported by: - The Belgian Science Policy (Belspo), through the Belgian Scientific Research Programme on the Antarctic and the “biodiversity.aq” network (SCAR-MarBIN/ANTABIF) The Royal Belgian Institute of Natural Sciences (RBINS), Brussels, Belgium The British Antarctic Survey (BAS), Cambridge, United Kingdom The Université Pierre et Marie Curie (UPMC), Paris, France The Australian Antarctic Division, Hobart, Australia The Scientific Steering Committee of CAML, Michael Stoddart (CAML Administrator) and Victoria Wadley (CAML Project Manager) Mapping coordination and design: Huw Griffiths (BAS, Cambridge) & Anton Van de Putte (RBINS, Brussels) Editorial assistance: Henri Robert, Xavier Loréa, Charlotte Havermans, Nicole Moortgat (RBINS, Brussels) Printed by: Altitude Design, Rue Saint Josse, 15, B-1210 Brussels, Belgium (www.altitude-design.be) Lay out: Sigrid Camus & Amélie Blaton (Altitude Design, Brussels). Cover design: Amélie Blaton (Altitude Design, Brussels) and the Editorial Team. Cover pictures: amphipod crustacean (Epimeria rubrieques De Broyer & Klages, 1991), image © T. Riehl, University of Hamburg; krill (Euphausia superba Dana, 1850), image © V. Siegel, Institute of Sea Fisheries, Hamburg; fish (Chaenocephalus sp.), image © C. d’Udekem d’Acoz, RBINS; emperor penguin (Aptenodytes forsteri G.R. Gray, 1844), image © C. d’Udekem d’Acoz, RBINS; Humpback whale (Megaptera novaeangliae (Borowski, 1781)), image © L. Kindermann, AWI. Online dynamic version : A dynamic online version of the Biogeographic Atlas will be available on the SCAR-MarBIN / AntaBIF portal : atlas.biodiversity.aq. Recommended citation: For the volume: De Broyer C., Koubbi P., Griffiths H.J., Raymond B., Udekem d’Acoz C. d’, Van de Putte A.P., Danis B., David B., Grant S., Gutt J., Held C., Hosie G., Huettmann F., Post A., Ropert-Coudert Y. (eds.), 2014. Biogeographic Atlas of the Southern Ocean. Scientific Committee on Antarctic Research, Cambridge, XII + 498 pp. For individual chapter: (e.g.) Crame A., 2014. Chapter 3.1. Evolutionary Setting. In: De Broyer C., Koubbi P., Griffiths H.J., Raymond B., Udekem d’Acoz C. d’, et al. (eds.). Biogeographic Atlas of the Southern Ocean. Scientific Committee on Antarctic Research, Cambridge, pp. xx-yy. ISBN: 978-0-948277-28-3. This publication is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License 2 Biogeographic Atlas of the Southern Ocean Palaeo-oceanography 3.3. Palaeo-Oceanography (Box) Rainer Gersonde Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research, Bremerhaven, Germany -30° 0° 30° E-LGM 14°C E-LGM 8°C STF 4°C GM -60° 60° PF SA F E-L SF E-LGM 0°C -90° -30° -45° -60° ? PF ? -120° 90° SAF ? 120° STF Palaeo-oceanography Map 1 Averaged Southern Ocean sea surface isotherms (°C) at the Last Glacial Maximum (LGM) (modified from Gersonde et al. 2005). Modern locations of the Polar Front (PF), Sub-Antarctic Front (SAF) and Sub-Tropical Front (STF) are shown for comparison (based on Belkin & Gordon 1996). The reconstruction relies on the estimation of austral summer sea surface temperatures from 107 sediment cores using transfer function techniques. The significance of individual estimates was ranked according to quality criteria concerning reliability of estimate and stratigraphic control (Gersonde et al. 2005). Signal carriers are diatom and radiolarian assemblages preserved in the sediment record. Closed points indicate locations with diatom-based, crosses with radiolarian-based reconstructions. The surface isotherms E-LGM 4°C, 8°C and 14°C stand for average locations of the Polar Front, Sub-Antarctic Front and Sub-Tropical Front during the EPILOG-LGM austral summer. EPILOG time slice placed between 23,000 and 19,000 cal. yr BP (modified from Gersonde et al. 2005). In the case of multiple sea surface temperatures (SST) estimates at any one location, only the highest quality estimate was considered for definition of the isotherm location. The E-LGM summer 0°C isotherm is close to the sporadic extent of summer sea ice during the LGM. The maximum extent of the EPILOG-LGM winter sea ice field coincides to some extent with the E-LGM 4°C isotherm. Both, LGM winter- and summer sea ice extent have been reconstructed based on a specific sea ice transfer function and additionally using diatom sea ice indicators. For more details on the sea ice reconstruction see Gersonde et al. (2005). Summer SSTs reveal greatest surface-water cooling in the area of the modern Sub-Antarctic Zone reaching a temperature decline of 4-6°C. As a result of the northward expansion of Antarctic cold waters by 5-10° in latitude and a relatively small displacement of the Sub-Tropical Front, thermal gradients were steepened during the last glacial in the northern zone of the Southern Ocean. The distribution of core locations available for the reconstruction shows that data coverage in the Pacific remains weak. A new expedition to this prominent sector of the Southern Ocean (Gersonde 2011) collected materials, which allow filling this gap in the very near future. References Belkin, I.M., Gordon, A.L.,1996. Southern Ocean fronts from the Greenwich meridian to Tasmania. Journal of Geophysical Research: Oceans 101 (C2), 3675–3696. Gersonde, R., Crosta, X., Abelmann, A., Armand, L. K., 2005. Sea surface temperature and sea ice distribution of the Southern Ocean at the EPILOG last glacial maximum — A circum-Antarctic view based on siliceous microfossil records. Quaternary Science Reviews, 24, 869–896. Gersonde, R., 2011. The Expedition of the Research Vessel “Polarstern” to the Polar South Pacific Sea in 2009/2010 (ANT-XXVI/2). Reports on Polar and Marine Research 632, 325 p. (http:// epic.awi.de) Further selected references on Southern Ocean palaeo-oceanography: Armand, L. K., Leventer, A., 2003. Palaeo sea ice distribution –reconstruction and palaeoclimatic significance. In: Thomas D.N., Dieckmann G.S. (eds.). Sea Ice: Physics,Chemistry and Biology. Oxford: Blackwell Science Ltd, pp. 333–372. Armand, L.K., Leventer, A., 2010. Palaeo sea ice distribution and reconstruction derived from the geological record. In: Thomas D.N., Dieckmann G.S. (eds.). Sea Ice: an Introduction to its Physics, Biology, Chemistry, and Geology, Second edition. Wiley-Blackwell. pp. 469-530. Armand, L.K., Crosta, X., Quéguiner, B., Mosseri, J., Garcia, N., 2008. Diatoms preserved in surface sediments of the northeastern Kerguelen Plateau. Deep-Sea Research II 55, 677–692 Armand, L.K., Crosta, X., Romero, O., Pichon, J.J., 2005. The biogeography of major diatom taxa in Southern Ocean sediments: 1. Sea ice related species. Palaeogeography, Palaeoclimatology, Palaeoecology 223, 93– 126. Crosta, X., 2011. Marine diatoms in polar and sub-polar environments and their application to Late Pleistocene paleoclimate reconstruction. IOP Conf. Series: Earth and Environmental Science 14, 1-18. doi:10.1088/1755-1315/14/1/012006 Crosta X., Koç, N., 2007. Diatoms: From micropaleontology to isotope geochemistry. In: Developments in Marine Geology, Volume 1. Elsevier B.V. Crosta, X., Romero, O., Armand, L.K., Pichon, J.J., 2005. The biogeography of major diatom taxa in Southern Ocean sediments: 2. Open ocean related species. Palaeogeography, Palaeoclimatology, Palaeoecology 223, 66– 92. Leventer, A., Crosta, X., Pike, J., 2010. Holocene marine diatom records of environmental change. In: Smol, J.P., Stoermer, E.F. (eds.). The diatoms: applications for the environmental and earth sciences. Cambridge University Press. pp. 401-423. Romero, O.E., Armand, L.K., Crosta, X., Pichon, J.J. 2005. The biogeography of major diatom taxa in Southern Ocean surface sediments: 3. Tropical/Subtropical species. Palaeogeography, Palaeoclimatology, Palaeoecology 223, 49– 65 Biogeographic Atlas of the Southern Ocean 43 THE BIOGEOGRAPHIC ATLAS OF THE SOUTHERN OCEAN Scope Biogeographic information is of fundamental importance for discovering marine biodiversity hotspots, detecting and understanding impacts of environmental changes, predicting future distributions, monitoring biodiversity, or supporting conservation and sustainable management strategies. The recent extensive exploration and assessment of biodiversity by the Census of Antarctic Marine Life (CAML), and the intense compilation and validation efforts of Southern Ocean biogeographic data by the SCAR Marine Biodiversity Information Network (SCAR-MarBIN / OBIS) provided a unique opportunity to assess and synthesise the current knowledge on Southern Ocean biogeography. The scope of the Biogeographic Atlas of the Southern Ocean is to present a concise synopsis of the present state of knowledge of the distributional patterns of the major benthic and pelagic taxa and of the key communities, in the light of biotic and abiotic factors operating within an evolutionary framework. Each chapter has been written by the most pertinent experts in their field, relying on vastly improved occurrence datasets from recent decades, as well as on new insights provided by molecular and phylogeographic approaches, and new methods of analysis, visualisation, modelling and prediction of biogeographic distributions. A dynamic online version of the Biogeographic Atlas will be hosted on www.biodiversity.aq. The Census of Antarctic Marine Life (CAML) CAML (www.caml.aq) was a 5-year project that aimed at assessing the nature, distribution and abundance of all living organisms of the Southern Ocean. In this time of environmental change, CAML provided a comprehensive baseline information on the Antarctic marine biodiversity as a sound benchmark against which future change can reliably be assessed. CAML was initiated in 2005 as the regional Antarctic project of the worldwide programme Census of Marine Life (2000-2010) and was the most important biology project of the International Polar Year 2007-2009. The SCAR Marine Biodiversity Information Network (SCAR-MarBIN) In close connection with CAML, SCAR-MarBIN (www.scarmarbin.be, integrated into www.biodiversity.aq) compiled and managed the historic, current and new information (i.a. generated by CAML) on Antarctic marine biodiversity by establishing and supporting a distributed system of interoperable databases, forming the Antarctic regional node of the Ocean Biogeographic Information System (OBIS, www.iobis.org), under the aegis of SCAR (Scientific Committee on Antarctic Research, www.scar.org). SCAR-MarBIN established a comprehensive register of Antarctic marine species and, with biodiversity.aq provided free access to more than 2.9 million Antarctic georeferenced biodiversity data, which allowed more than 60 million downloads. The Editorial Team Claude DE BROYER is a marine biologist at the Royal Belgian Institute of Natural Sciences in Brussels. His research interests cover structural and ecofunctional biodiversity and biogeography of crustaceans, and polar and deep sea benthic ecology. Active promoter of CAML and ANDEEP, he is the initiator of the SCAR Marine Biodiversity Information Network (SCAR-MarBIN). He took part to 19 polar expeditions. Philippe KOUBBI is professor at the University Pierre et Marie Curie (Paris, France) and a specialist in Antarctic fish ecology and biogeography. He is the Principal Investigator of projects supported by IPEV, the French Polar Institute. As a French representative to the CCAMLR Scientific Committee, his main input is on the proposal of Marine Protected Areas. His other field of research is on the ecoregionalisation of the high seas. Huw GRIFFITHS is a marine Biogeographer at the British Antarctic Survey. He created and manages SOMBASE, the Southern Ocean Mollusc Database. His interests include large-scale biogeographic and ecological patterns in space and time. His focus has been on molluscs, bryozoans, sponges and pycnogonids as model groups to investigate trends at high southern latitudes. Ben RAYMOND is a computational ecologist and exploratory data analyst, working across a variety of Southern Ocean, Antarctic, and wider research projects. His areas of interest include ecosystem modelling, regionalisation and marine protected area selection, risk assessment, animal tracking, seabird ecology, complex systems, and remote sensed data analyses. Cédric d’UDEKEM d’ACOZ is a research scientist at the Royal Belgian Institute of Natural Sciences, Brussels. His main research interests are systematics of amphipod crustaceans, especially of polar species and taxonomy of decapod crustaceans. He took part to 2 scientific expeditions to Antarctica on board of the Polarstern and to several sampling campaigns in Norway and Svalbard. Anton VAN DE PUTTE works at the Royal Belgian Institute for Natural Sciences (Brussels, Belgium). He is an expert in the ecology and evolution of Antarctic fish and is currently the Science Officer for the Antarctic Biodiveristy Portal www. biodiversity.aq. This portal provides free and open access to Antarctic Marine and terrestrial biodiversity of the Antarctic and the Southern Ocean. Bruno DANIS is an Associate Professor at the Université Libre de Bruxelles, where his research focuses on polar biodiversity. Former coordinator of the scarmarbin. be and antabif.be projects, he is a leading member of several international committees, such as OBIS or the SCAR Expert Group on Antarctic Biodiversity Informatics. He has published papers in various fields, including ecotoxicology, physiology, biodiversity informatics, polar biodiversity or information science. Bruno DAVID is CNRS director of research at the laboratory BIOGÉOSCIENCES, University of Burgundy. His works focus on evolution of living forms, with and more specifically on sea urchins. He authored a book and edited an extensive database on Antarctic echinoids. He is currently President of the scientific council of the Muséum National d’Histoire Naturelle (Paris), and Deputy Director at the CNRS Institute for Ecology and Environment. Susie GRANT is a marine biogeographer at the British Antarctic Survey. Her work is focused on the design and implementation of marine protected areas, particularly through the use of biogeographic information in systematic conservation planning. Julian GUTT is a marine ecologist at the Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research, Bremerhaven, and professor at the Oldenburg University, Germany. He participated in 13 scientific expeditions to the Antarctic and was twice chief scientist on board Polarstern. He is member of the SCAR committees ACCE and AnT-ERA (as chief officer). Main focii of his work are: biodiversity, ecosystem functioning and services, response of marine systems to climate change, non-invasive technologies, and outreach. Christoph HELD is a Senior Research Scientist at the Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research, Bremerhaven. He is a specialist in molecular systematics and phylogeography of Antarctic crustaceans, especially isopods. Graham HOSIE is Principal Research Scientist in zooplankton ecology at the Australian Antarctic Division. He founded the SCAR Southern Ocean Continuous Plankton Recorder Survey and is the Chief Officer of the SCAR Life Sciences Standing Scientific Group. His research interests include the ecology and biogeography of plankton species and communities, notably their response to environmental changes. He has participated in 17 marine science voyages to Antarctica. Falk HUETTMANN is a ‘digital naturalist’ he works on three poles ( Arctic, Antarctic and Hindu-Kush Himalaya) and elsewhere (marine, terrestrial and atmosphere). He is based with the university of Alaska-Fairbank (UAF) and focuses primarily on effective conservation questions engaging predictions and open access data. Alexandra POST is a marine geoscientist, with expertise in benthic habitat mapping, sedimentology and geomorphic characterisation of the seafloor. She has worked at Geoscience Australia since 2002, with a primary focus on understanding seafloor processes and habitats on the East Antarctic margin. Most recently she has led work to understand the biophysical environment beneath the Amery Ice Shelf, and to characterise the habitats on the George V Shelf and slope following the successful CAML voyages in that region. Yan ROPERT COUDERT spent 10 years at the Japanese National Institute of Polar Research, where he graduated as a Doctor in Polar Sciences in 2001. Since 2007, he is a permanent researcher at the CNRS in France and the director of a polar research programme (since 2011) that examines the ecological response of Adélie penguins to environmental changes. He is also the secretary of the Expert Group on Birds and Marine Mammals and of the Life Science Group of the Scientific Committee on Antarctic Research. AnT-ERA